275

Tropical Bryology 8: 275-280, 1993

Notes on the amphipacific relations of Hawaiian

S. Stenroos Department of Botany, P.O. Box 47, FIN-00014 Helsinki, Finland

C.W. Smith Department of Botany of the University of Hawaii at Manoa, Honolulu, Hawaii 96822, USA

Abstract. The total number of currently accepted of Cladoniaceae in the Hawaiian Islands is 22. Several taxonomic problems still exist, however. The effects of isolation are clear among Cladoniaceae. Endemism is high (c. 40%); and, the number of species low. The species must have reached the archipelago via long-distance trans-oceanic dispersal, probably aided by the abundant production of propagules, such as soredia and microsquamules. Although most of the species found in Hawaii are widely distributed, the Hawaiian Cladoniaceae show slight affinities to those of E and SE Asia. polyphylla Mont. & v.d. Bosch is an older name for C. fruticulosa Krempelh., and is lectotypified from authentic material. C. leprosula H. Magn. is included in C. ochrochlora Flörke.

True oceanic islands, like the Hawaiian Islands, islands (Hawaii, Kahoolawe, Maui, Lanai, Mo- have never had any connection or association lokai, Oahu, Kauai, and Niihau), the largest of with continental land during geographic diversi- which - and still with volcanic activity - being fication that has been going on through geologi- Hawaii (c. 1560 sq km). cal time. These islands are volcanoes arising from the ocean floor. The Hawaiian Islands are Major works on Hawaiian , including shield volcanoes that are formed where tectonic Cladoniaceae, are those of des Abbayes (1947), plates move over a “hot spot” and magma wells Magnusson (1956, who summarizes exhaustive- up to the surface (e.g. Press & Siever 1982, ly the earlier information), Klement (1966) and Fosberg 1991). The estimated age of the Hawai- Smith (1981). Some ecologial investigations ian archipelago is c. 50 million years. Today the including Cladoniaceae have also been reported nearest continent is North America (c. 3800 km (e.g. Kappen & Smith 1980). to the northeast). Japan is c. 5400 km to the west, and the closest oceanic atolls are c. 800 km The main purpose of the present paper is to distant. The archipelago comprises eight major update information on Hawaiian species of the 276

Cladoniaceae, including their . In Magnusson 1956), C. pyxidata (L.) Fr., C. addition, we show the importance of long-distan- scabriuscula (Delise) Nyl., C. sphacelata Vainio, ce dispersal that seems to be surprisingly effective and C. subradiata (Vainio) Sandst. (as C. among Cladoniaceae in many areas, especially fimbriata var. balfourii (Crombie) Vainio). Of in the Southern Hemisphere where land is the reindeer lichens, Cladina rangiferina (L.) scattered (see e.g. Galloway 1991, Stenroos 1993). Nyl. and C. mitis (Sandst.) Hustich were definitely excluded from the Hawaiian flora by Ahti (1961). We emphasize that the material used for this study is not complete. Some specimens could not In addition, the following taxa reported from the be determined to any known species, but on the Islands are reduced to synonymy, further redu- other hand we were reluctant to describe new cing the total number of accepted species present: taxa based on sparse material. Others have not Cladonia congregata H. Magn. (= C. didyma been relocated, some destroyed in World War II. (Fée) Vainio, see des Abbayes 1966, Stenroos The results are mainly based on specimens in H, 1986a), C. fauriei des Abb. (= C. kauaiensis HAW and TUR (which include the collection Merrill, see Stenroos 1986a), C. leprosula H. made by L. & Y. Mäkinen in 1966), and on field Magn. (= C. ochrochlora Flörke, T. Ahti pers. work of both authors. Several specimens were comm.), C. subsquamosa (Nyl.) Crombie (= C. analysed by standardized thin-layer chromato- squamosa Hoffm., see Huovinen & Ahti 1988), graphic methods (Culberson 1972, White & James and C. vulcanica Zoll. & Moritzi (= C. didyma 1985). (Fée) Vainio, see Huovinen et al. 1989). C. polyphylla Mont. & v.d. Bosch (as C. pityrea *polyphylla (Mont. & V.D. Bosch) des Abb. in Results and discussion Magnusson 1956) was found here to be conspe- cific with C. fruticulosa Krempelh. However, Altogether 22 species of Cladina and Cladonia although C. polyphylla (lectotype, here selected: are recorded in this study (Table 1). The number Java, Junghuhn, PC-Hue; syntypes, L, PC-Hue) of species reported from the area by earlier is an older name for this taxon, it is not accepted authors is almost twice as high (see Magnusson here because C. fruticulosa shall be proposed for 1956, who recognized 37 species). All previously conservation according to the new Art. 14.1 in recorded species have been assigned to currently the Botanical Code as approved in the XV Inter- extant species in the islands; none are extinct. national Botanical Congress in Yokohama in 1993. The following species reported from Hawaii by earlier authors are excluded from the flora becau- Finally, in addition to Cladina magnussonii (Ahti) se in light of the material examined and the Ahti, which was reported by Ahti (1961), the knowledge of the distribution of the species following taxa are recorded here as additions to elsewhere they are unlikely to occur in the Magnusson’s (1956) list of Cladoniaceae of the Hawaiian Islands, or their presence needs Hawaiian Islands: Cladonia cf. corniculata Ahti confirmation: aggregata (Sw.) Nyl., & Kashiw., C. merochlorophaea Asah. var. Cladonia chlorophaea (Flörke) Sprengel (as C. merochlorophaea, C. pleurota (Flörke) Schae- pyxidata var. chlorophaea Flörke in Magnusson rer s.lat., and C. poeciloclada des Abb. One 1956), C. coniocraea (Flörke) Sprengel, C. specimen (Kauai, Hanalei, Bowler 1517, H), corallifera (Kunze) Nyl., C. digitata (L.) Hoffm., with farinose soredia, imperfectly scyphose C. erythrosperma Vainio, C. fimbriata (L.) Fr., podetia, and fumarprotocetraric acid, probably C. furcata (Hudson) Schrader, C. gracilis (L.) belongs to an undescribed species (“sp. 1” in Willd., C. granulosa (Vainio) Ahti (as C. Table 1). Specimens (Kauai, Kokee, L. & Y. subsquamosa f. granulosa Vainio in Magnusson Mäkinen 66-2031, TUR; Oahu, Waianae Range, 1956), C. grayi Merr. ex Sandst., C. macilenta L. & Y. Mäkinen 66-1627, TUR; Hawaii, Hoffm., C. mexicana Vainio, C. phyllophora Volcanoes Nat. Park, Stenroos 3918, H) Hoffm. (as C. degenerans (Flörke) Sprengel in determined here as “sp. 2” are related to C. 277

Table 1. List of Hawaiian Cladoniaceae and their major secondary substances. BAR=barbatic acid (incl. 4-O-demethylbarbatic acid), DID=didymic acid, FUM-complex=fumarprotocetraric and protocetraric acids and Cph-2 (Cph- 1 present or absent), HSEK=homosekikaic acid, IUSN=isousnic acid, MCRY=4'-O- methylcryptochlorophaeic acid, MER=merochlorophaeic acid, PER=perlatolic acid, PSO=psoromic acid, SQU=squamatic acid, THA=thamnolic acid, USN=usnic acid. ———————————————————————————————————— —————— Species Major secondary substances ———————————————————————————————————— —————— Cladina leiodea (H. Magn.) Ahti USN, FUM-complex magnussonii (Ahti) Ahti ATR, FUM-complex skottsbergii (H. Magn.) Follm. USN, PER f. fuscescens (Ahti) Ahti PER Cladonia angustata Nyl. USN, BAR cf. cenotea (Ach.) Schaerer SQU cf. corniculata Ahti & Kashiw. FUM-complex didyma (Fée) Vainio THA, DID or BAR farinacea (Vainio) A. Evans FUM-complex fruticulosa Krempelh. PSO or FUM-complex kauaiensis G.K. Merrill ex H.Magn. & Zahlbr. USN, THA mauiensis H. Magn. BAR, DID merochlorophaea Asah. var. merochlorophaea MCRY, MER, FUM-complex oceanica (Vainio) Zahlbr. USN, BAR, DID ochrochlora Flörke FUM-complex pleurota (Flörke) Schaerer USN, IUSN, ZEO poeciloclada des Abb. FUM-complex ramulosa (With.) Laundon FUM-complex, HSEK, ATR (incl. C. adspersa Mont. & Bosch) solitaria H. Magn. USN squamosa Hoffm. SQU subsquamosa Krempelh. FUM-complex ramulosa and C. phyllopoda (Vainio) Stenroos as well. However, this complex still needs a (Stenroos 1988) but differ from them by their thorough revision. For some of the more poorly very slender, more branched, granulose, and known taxa of those accepted here, the reader is partly escyphose podetia. Specimens generally referred to some previous papers: Cladonia referred to as C. scabriuscula or C. farinacea are angustata Nyl. (Stenroos 1986a), C. fruticulosa here included in C. farinacea s.lat. The present Krempelh. (as C. decipiens des Abb. in material is often richly squamulose (in contrast Magnusson 1956, see Stenroos 1988), C. to C. farinacea s.str. in South America) and the mauiensis H. Magn. (Stenroos 1986a), C. podetial squamules and parts of podetial surface oceanica (Vainio) Zahlbr. (Stenroos 1986a), and are farinose-sorediate (contrary to C. scabriuscula C. solitaria H. Magn. (Stenroos 1986b). s.str.). Similar material is found in Eastern Asia 278

The effects of prolonged isolation of the Hawai- have weak affinities with Central and South ian Islands are clear in taxa of the Cladoniaceae. America (Smith 1993, 1994). The degree of endemism in lichens is high, viz. c. 40% (parmelioid lichens 8%, alectorioid li- Successful dispersal is not only a problem of how chens 0%, umbilicarioid lichens 0%, lichens to reach an area but also of establishment once total 38% (see Magnusson 1956), mosses 46% there. Asexual propagules, produced abundantly (see Hoe 1979), ferns & fern allies 70% (C. H. in most Cladoniaceae, have both bionts present. Lamoureux, pers. comm.), and angiosperms Yet most species are slow in their development 95%), and the number of species is relatively and need stable conditions without much compe- low. Because of the isolation of the archipelago, tition or disturbance. Both competition and di- taxa in the Cladoniaceae must have reached the sturbance are encountered on Hawaii, where area via long-distance trans-oceanic dispersal luxurious vascular vegetation with many agres- or, in the case of endemics, evolved on the sive alien plants has taken over many areas. islands. Cladoniaceae are also absent from large arable and cultivated areas and areas under grazing Long-distance dispersal is apparently aided by (mainly cattle). Feral pig activity in natural the abundant production of lichen propagules forests results in a dramatic decrease in cladoni- such as soredia and microsquamules which are oid species except on rotting logs which are very common in the Cladoniaceae. The priority rarely disturbed. Volcanic activity is extremely of vegetative dispersal is supported by the fact destructive: lava flows may bury large areas, and that spore-production seems to be poor, at least in bare volcanic rock and heat certainly set limits to many present-day Cladoniaceae, even if the the growth of many Cladoniaceae. On the other species is frequently fertile. hand, ash fields and lava flows in wet areas provide excellent habitats for some species. The Ancestors of the esorediate and esquamulose invasion of alien bunchgrasses this century has endemics of Cladina as well as of Clado- not only crowded out cladonioid lichens from the nia sect. Unciales (viz. C. solitaria) may have many areas of exposed lava, but also dramatical- reached the islands by means of easily disintegra- ly increased the incidence and severity of wildfi- ting thallus fragments. Most of the other species res (Smith and Tunison 1992). The immediately possess either farinose or granulose soredia (C. adjacent regions of thermal areas also provide angustata, C. cf. cenotea, C. cf. corniculata, C. habitats e.g. for the endemic C. oceanica (Kap- fruticulosa, C. merochlorophaea, C. ochrochlo- pen & Smith 1980, Smith 1981). The Islands, ra, C. pleurota s.lat., C. poeciloclada, C. therefore, provide a wide array of habitats though farinacea s.lat., and the endemics C. mauiensis, many of them are now inimical to lichens. C. oceanica, and C. sp. 1) or microsquamules or granules (C. didyma, C. ramulosa, C. squamosa, Of the endemics, Cladina leiodea, C. skottsber- and C. sp. 2). The endemic C. kauaiensis produ- gii, Cladonia oceanica, and C. sp. 2 are distribu- ces relatively large, attached granules but those ted on most islands, whereas the others seem to of its ancestors may have been more suitable for be confined to a smaller area: Cladina long-distance dispersal. magnussonii (Maui), Cladonia kauaiensis (Kauai), C. mauiensis (Maui), C. solitaria (Maui), Most of the species which also occur outside C. sp. 1 (Kauai); Cladina magnussonii and Hawaii, have a relatively wide total range (C. Cladonia solitaria are actually confined to a very didyma, C. merochlorophaea, C. ochrochlora, small area at the summit of W. Maui. C. pleurota, C. ramulosa, C. farinacea s. lat., and C. squamosa). However, as with the phane- rogams (Carlquist 1970) the Hawaiian Clado- niaceae flora seems to show some affinities with that of E and SE Asia (Fig. 1). On the other hand, the alectorioid and parmelioid lichens of Hawaii Acknowledgements

280

Hoe, W. J. 1979. The phytogeographical relationships of Hawaiian mosses. Ph. D. Thesis, Univ. Hawaii, Honolulu. Huovinen, K. & Ahti, T. 1988. The composition and contents of aromatic lichen substances in Cladonia section Perviae. Ann. Bot. Fennici 25:371-383. Huovinen, K., Ahti, T. & S. Stenroos. 1989. The composition and contents of aromatic lichen substances in Cladonia, section Cocciferae. Ann. Bot. Fennici 26: 133-148. Kappen, L. & C. W. Smith. 1980. Heat tolerance of two Cladonia species and Campylopus praemorsus in a hot steam vent area of Hawaii. Oecologia (Berl.) 47: 184-189. Klement, O. 1966. Zur Kenntnis der Flechtenflora und - vegetation des Hawaii-Archipels. I. Lanai. Nova Hedwigia 11: 243-283. Magnusson, A. H. 1956(‘1955’). A catalogue of the Hawaiian lichens. Arkiv Bot., ser. 2, 3: 223-402. Press, F. & R. Siever. 1982. Earth. 613 pp. W.H. Freeman & Co., San Francisco. Smith, C. W. 1981. Bryophytes and lichens of the Puhimau geothermal area, Hawaii Volcanoes National Park. Bryologist 84: 457-466. Smith, C. W. 1991. Impact of alien plants on Hawaii’s native biota. In Stone C.P. & J.M. Scott (eds.), Hawaii’s terrestrial ecosystems: Preservation and management. Cooperative Nat. Parks Resources Studies Unit, Univ. Hawaii, Honolulu. Smith, C.W. 1993. Notes on Hawaiian parmeloid lichens. Bryologist 96: 326-332. Smith, C.W. 1994. Hawaiian Alectorioid lichens. Bryologist (in press). Smith, C. W. & J. T. Tunison. 1992. Fire and alien plants in Hawai‘i: Research and management implications for native ecosystems. In Stone, C. P., C. W. Smith & J.T. Tunison (eds.), Alien plant invasions in native ecosystems of Hawai‘i: Manage- ment and research. Cooperative Nat. Parks Resources Studies Unit, Univ. Hawaii, Honolulu. Stenroos, S. 1986a. The family Cladoniaceae in Melanesia. 2. Cladonia section Cocciferae. Ann. Bot. Fennici 23: 239-250. Stenroos, S. 1986b. The family Cladoniacae in Melanesia. 1. Cladonia sect. Unciales. Ann. Bot. Fennici 23: 161-164. Stenroos, S. 1988. The family Cladoniaceae in Melanesia. 3. Cladonia sections Helopodium, Perviae and Cladonia. Ann. Bot. Fennici 25: 117-148. Stenroos, S. 1993. Taxonomy and distribution of the lichen family Cladoniaceae in the Antarctic and peri-Antarctic re- gions. Crypt. Bot. (in press). White, J. & P. W. James. 1985. A new guide to microchemical techniques for the identification of lichen substances. British Lich. Soc. Bull. 57 (Suppl.): 1-41.